MADRON˜ O, Vol. 54, No. 4, pp. 269–274, 2007 LONG-TERM GROWTH AND PERSISTENCE OF BLUE OAK (QUERCUS DOUGLASII) SEEDLINGS IN A CALIFORNIA OAK SAVANNA WALTER D. KOENIG Hastings Reservation and Museum of Vertebrate Zoology, University of California Berkeley, 38601 E. Carmel Valley Road, Carmel Valley, CA 93924 [email protected] JOHANNES M. H. KNOPS School of Biological Sciences, University of Nebraska, 348 Manter Hall, Lincoln, NE 68588 ABSTRACT We report on growth and survivorship of two cohorts of blue oaks Quercus douglasii Hook. & Arn. (Fagaceae) monitored at Hastings Reservation in Monterey County, California, the first growing in an unprotected old field and measured as seedlings in 1965 and the second planted as acorns in 1985 in several sites differing in their degree of protection from grazing. Growth of all individuals was extremely slow: among those surviving in the first cohort, mean (6 SD) height in 2006 was only 76.7 6 45.0 cm for an average growth rate of 1.8 cm yr21, and only one of the original 73 oaks had grown taller than 1.5 m while one was still a seedling 28 cm in height 41 yr after being first marked. Of the second cohort, mean height 21 yr after planting was 54.3 6 31.4 cm. None of these latter individuals had grown out of the sapling stage while 25% were still seedlings , 30 cm in height. Growth of this second cohort was significantly greater when protected from grazing and when growing in the open rather than in the shade. Although growth was slow, survivorship of oaks first measured in 1965 was high, indicting that individuals can live for decades despite significant grazing pressure. Our results confirm the difficulties of inferring age from size of blue oaks, since individuals just achieving the height at which they are typically cored may be 50 or more years old. They also indicate that regeneration, although very slow, can occur in open oak savannas in California despite significant grazing pressure. Whether the observed amount of regeneration is sufficient for long-term sustainability will require continued monitoring and modeling of oak demography. Key Words: blue oak, Quercus douglasii, regeneration, seedling growth, survivorship. The health and status of California’s vast oak natural seedling recruits as part of studies going woodlands is one of the more vexing questions back to 1965. Our results indicate that the currently facing the state’s ecologists and range- relationship between size and age in this species land managers. Are they declining, and if so, is it may be even more problematical than previously due to grazing, competition from exotic grasses, suspected. They also confirm the remarkable fire suppression, climate change, or some combi- degree to which blue oak saplings can cling to nation of factors (Griffin 1981; Muick and what appears to be a precarious existence over Bartolome 1987; Gordon and Rice 2000; Kuep- many years until such time that they are able to pers et al. 2005; Tyler et al. 2006)? In the case of achieve sufficient height to escape browsing. the blue oak Quercus douglasii Hook. & Arn. (Fagaceae), a species dominating nearly 2 3 METHODS 106 ha in the state (Standiford 2002), the evidence is mixed, as demographic surveys almost uni- The study was conducted at Hastings Reser- formly reveal limited recruitment but long-term vation, Monterey County, California, established studies have generally shown no decline in tree in 1937. Thus, although cattle grazing, clearing, density (Tyler et al. 2006). and various agricultural activities occurred his- Resolving this problem lies in obtaining more torically, no such disturbances took place during extensive data on the demography of the species the time period covered by this study. Hastings is in question. Unfortunately this has not proved to located approximately 50 km from the coast and be easy, since individuals can live hundreds of averages 53.3 cm of rain year21 (mean of 67 yr years and are logistically difficult to age, making between 1 July 1939–30 June 1940 and 1 July it virtually impossible to estimate when prior 2005–30 June 2006; records from Reserve head- regeneration events occurred, much less the quarters). Individual oaks followed were either extent to which such events are episodic and planted or monitored in two areas of the Reserve. dependent on particular ecological circumstances. The first, North Field, was an old field cleared Here we report on the persistence and growth around 1900 and used until 1937 for hay and of blue oaks planted as acorns or first marked as grapes. North Field was left open the entire time 270 MADRON˜ O [Vol. 54 and was thus regularly grazed by mule deer cover of a mature blue oak within the plot; (Odocoileus hemionus) as well as smaller mam- seedlings in both these blocks were also protected mals including gophers and mice. The second by wire mesh baskets 40 cm in height. For area (Arnold Road Flat) was within a large analysis, we divided individuals into those that exclosure built in 1983 for a separate study and were not protected (block 1) vs. protected (blocks thus was not grazed by large herbivores during 2–4)(variable ‘‘protection’’), and those that were the study, but was open to smaller mammals. growing in the open (blocks 2 and 3) vs. those Two sets of oaks were examined. The first, that were shaded (blocks 1 and 4)(variable called the ‘‘White’’ cohort, consisted of 73 ‘‘shade’’). naturally-recruiting blue oak seedlings in North We measured the maximum height, maximum Field that were tagged and measured (height width, and basal diameter at 5 cm above ground only) in April 1965 by K. L. White. Three of the on 5 October 2006 of all individuals we could seedlings were new in spring 1965. All others were unambiguously identify as having been from the pre-1965 seedlings when tagged. Their ages were original set of acorns based on remaining wooden not known, but all were short (mean 6 SD 5 6.5 stakes and their location within the original grid 6 3.2 cm, range 2–18 cm) and were presumably on which acorns were originally planted (Fig. 1). thought to have been from acorns produced in All individuals were thus known to be 21 yr old fall of 1963 that sprouted in spring 1964. It at the time they were measured. No data from was not possible to identify the original 1965 vs. prior years was available. We used the two pre-1965 seedlings, and thus all individuals in the categories of ‘‘protection’’ and ‘‘shade’’ to White cohort are assumed to have been at least quantify the effects of these variables on growth 42 yr old in 2006. of the saplings using general linear models. Subsequent to marking, seedlings and saplings Survivorship could not be measured in these were monitored and measured by J. R. Griffin on oaks, as we could not be certain how many had 10 occasions (June 1969, January 1976, Septem- been planted initially. ber 1979, January 1984, June 1986, June 1987, Following Phillips et al. (1997), we classify August 1988, June 1989, June 1990, and Febru- individuals as ‘‘seedlings’’ (,30 cm in height), ary 1991). With some exceptions, at each ‘‘saplings’’ (30–150 cm in height), ‘‘poles’’ (150– sampling period the height of tallest sprout from 300 cm in height), and ‘‘adults’’ (taller than 3 m). each remaining seedling was measured, while Significant browsing by deer, where present, is starting in 1988, the greatest width of each expected up until individuals reach the pole size seedling was measured. Finally, on 5 October class (McCreary 2001). Values presented are 2006, we examined all individuals whose identity means 6 SD. we could be confident of and measured their height, width, and the basal diameter of stems . RESULTS 0.5 cm at 5 cm above the ground. When multiple stems were present, they were combined to yield White Cohort a single value for the overall diameter that matched the total basal area of all stems. Size Of the original 73 seedlings marked in 1965, J. and growth rates are based on the 14 individuals R. Griffin found 64 (88%)inJune1969and18 we identified and measured in 2006. (24.7%) in February 1991, while we successfully The second set of oaks used in the study, called located 14 (19.2%) in 2006 (Fig. 2). These values the ‘‘Menke’’ cohort, consisted of individuals represent minimum survivorship of the original from a study of water relations of California oaks seedlings, since other seedlings and saplings were by J. W. Menke initiated in 1985. Blue oak acorns present in the plot and it is possible that some lost were planted in several 5 3 5 blocks varying in their tags during the course of the study and their degree of protection from grazing and could no longer be identified. openness. Block 1 (not protected; shaded) was Growth of the seedlings varied considerably, planted adjacent to several blue oak trees near with individuals at the end of the study an North Field, but otherwise not protected from average of 76.7 6 45.0 cm (range 28 to 200 cm) grazing in any way. Block 2 (protected; open), in height (Fig. 3), 90.6 6 43.8 cm (range 13 to also near North Field, consisted of acorns 174 cm) in width, and 4.3 6 2.0 cm (range 0.7 to planted in the open, but protected by wire mesh 7.7 cm) in basal diameter. The rate of growth was baskets up to 40 cm in height that were opened , 1cmyr21 in height during the first three up in 1992 to allow free growth of saplings that decades (Fig.